See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/292678428 Cichlids of the Banc d'Arguin National Park, Mauritania: Insight into the diversity of the genus Coptodon Article in Journal of Fish Biology · February 2016 DOI: 10.1111/jfb.12899 CITATIONS READS 0 193 11 authors, including: Andreas R Dunz Antoine Pariselle Zoologische Staatssammlung München Institute of Research for Development 6 PUBLICATIONS 68 CITATIONS 122 PUBLICATIONS 915 CITATIONS SEE PROFILE SEE PROFILE Camilo Carneiro Jean-Dominique Durand Aveiro University & Iceland University Institute of Research for Development 7 PUBLICATIONS 6 CITATIONS 134 PUBLICATIONS 2,135 CITATIONS SEE PROFILE SEE PROFILE Some of the authors of this publication are also working on these related projects: IRD / LIPI PARI project View project systematic of fish parasites View project All content following this page was uploaded by Jean-Dominique Durand on 23 February 2016. The user has requested enhancement of the downloaded file. All in-text references underlined in blue are added to the original document and are linked to publications on ResearchGate, letting you access and read them immediately. Journal of Fish Biology (2016) doi:10.1111/jfb.12899, available online at wileyonlinelibrary.com Cichlids of the Banc d’Arguin National Park, Mauritania: insight into the diversity of the genus Coptodon N. G. Kide*, A. Dunz†, J.-F. Agnèse‡, J. Dilyte§, A. Pariselle‡‖, C. Carneiro¶, E. Correia¶, J. C. Brito§, L. O. Yarba¶, Y. Kone** and J.-D. Durand§§ ‡‡ *Université Abdel Maleck Essaâdi - Faculté des Sciences, Département de Biologie, Laboratoire Biologie Appliqué et Pathologie, Tétouan, Morocco, †Bavarian State Collection of Zoology, Department of Ichthyology, Münchhausenstr. 21, 81247 München, Germany, ‡Institut de Recherche pour le Développement (IRD), Université Montpellier, UMR 226, Institut des Sciences de l’Evolution de Montpellier, Place Eugène Bataillon, CC 65, F-34095, Montpellier cedex 5, France, §CIBIO/InBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos da Universidade do Porto Instituto de Ciências Agrárias de Vairão, R. Padre Armando Quintas, 4485-661 Vairão, Portugal, ¶Observatoire du Parc National du Banc d’Arguin, Nouakchott, Mauritania, ‖IRD, ISE-M, BP 1857 Yaoundé, Cameroon, **Faculté des Sciences de l’Université de Nouakchott, Nouakchott, Mauritania and §§Institut de recherche pour le développement (IRD), Université Montpellier, UMR MARBEC, Place Eugène Bataillon, CC 93, F-34095 Montpellier cedex 5, France (Received 28 May 2015, Accepted 17 December 2015) To determine the species diversity of cichlids in the Banc d’Arguin National Park (PNBA) and their phylogenetic relationships with other species in West Africa, a morphometric and meristic and molecular phylogenetic study was conducted. Both approaches not only confirm the presence of Sarotherodon melanotheron in PNBA but also demonstrate the presence of a second species from the genus Coptodon. While morphometric characteristics match the description of the Guinean tilapia Coptodon guineensis, phylogenetic reconstructions based on three mitochondrial and one nuclear DNA fragment demonstrate that C. guineensis is paraphyletic over its range. Because different lineages of C. guineensis are allopatric, the distribution of C. guineensis should be restricted to Ghana and Côte d’Ivoire. The many other lineages of this species should be considered as C. sp. aff. guineensis. © 2016 The Fisheries Society of the British Isles Key words: Cichlidae; cryptic species; phylogeny; West Africa. INTRODUCTION Cichlidae is one of the most speciose families of vertebrates, found in South and Central America, Madagascar, Africa, The Middle East, India and Sri Lanka (Kocher, 2004; Salzburger & Meyer, 2004). As with many other families of fishes, cichlid taxonomy is debated because systematics have been profoundly questioned and secondarily revised ‡‡Author to whom correspondence should be addressed. Tel.: + 84 1 223747334; email: Jean-Dominique. [email protected] 1 © 2016 The Fisheries Society of the British Isles 2 N. G. KIDE ET AL. with information from molecular phylogenetics (Chakrabarty, 2010; Dunz, 2012). Global cichlid diversity consists of major lineages with phylogenetic relationships consistent with Gondwanan break-up (Zardoya et al., 1996; Streelman et al., 1998; Sparks & Smith, 2004; McMahan et al., 2013). All African cichlid diversity belongs to the subfamily Pseudocrenilabrinae Fowler 1934 and includes the haplotilapiine formerly referred to as ‘Tilapia’ (Schwarzer et al., 2009; Dunz & Schliewen, 2013). The most recent comprehensive molecular phylogenetic review demonstrated that the haplotilapiine lineage is monophyletic and consists of 10 clades, with all East African cichlid tribes being recovered in a single clade (Dunz & Schliewen, 2013). Considering the molecular phylogeny and some diagnostic morphometric characteristics, Dunz & Schliewen (2013) proposed a novel generic and supra-generic classification of haplotilapiine. In this new classification, the tribes Oreochromini and Coptodonini harbour most of the genus and species diversity of West African cichlids. The extent of species diversity, however, remains to be evaluated because many new species are still being described (Dunz & Schliewen, 2010a, b; Dunz, 2012). Similar to the East African Great Lakes, Cameroon’s craters lakes in West Africa have been a focus of much study because they host species flocks (Trewavas et al., 1972; Stiassny et al., 1992; Schliewen et al., 1994; Schliewen & Klee, 2004; Neumann et al., 2011; Martin, 2012; Lamboj, 2014). In comparison, the African cichlid diversity outside of these regions appears to be extremely poor (Teugels & Thys van den Audenaerde, 2003). Only two species, the blackchin tilapia Sarotherodon melanotheron Rüppell 1852 and the Guinean tilapia Coptodon guineensis (Günther 1862), have been described so far in near-shore African marine environments, the latter being limited to Senegalese shores according to Stiassny et al. (2008). Some authors, however, reported the presence of C. guineensis in northern Moroccan localities: the Sebkhet Imlily, situated at c. 200 km from the border with Mauritania (Qninba et al., 2009), and 700 km to the North in rock pools (locally known as gueltas) along the Aabar wadi, tributary of Chbeyka wadi, Tan Tan province (south-western Morocco; Qninba et al., 2012). Similarly, while only one cichlid species is known in the Banc d’Arguin National Park (PNBA) in the north of Mauritania (Sevrin-Reyssac & Richer de Forges, 1985), Imraguen (the name given to local PNBA fishermen) distinguish two cichlids based on body colouration: ablack phenotype named Toumvertel kahla and a yellow phenotype named Toumvertel safra. Recent unpublished studies on cichlids targeted by fisheries at the PNBA also reported the presence of different phenotypes among the cichlids landed. All these independent observations call for further investigation to better describe the distribution range of cichlids in the Saharan area. Furthermore, because the taxonomic status of cichlids with widespread species has been controversial, such as for C. guineensis, the redbelly tilapia Coptodon zillii (Gervais 1848) and the redbreast tilapia Coptodon rendalli (Boulenger 1897) (for the list of synonymized species, see Eschmeyer, 2015), it is worthwhile to investigate the genetic status of these species further. In the past, genetic investigations of S. melanotheron demonstrated that this nominal species consisted of at least two species (and three sub-species) over its distribution from Congo to Mauritania (Falk et al., 1999, 2003). In this context, the first objective of this study was to investigate the phenotypic and genetic diversity of cichlids in the PNBA waters, by analysing their morphological and genetic variability, in order to clarify their taxonomic status. Secondly, the diversity of the tribe Coptodonini, and more specifically of some of their widespread species such as C. guineensis, C. rendalli or C. zillii, was evaluated by analysing several samples © 2016 The Fisheries Society of the British Isles, Journal of Fish Biology 2016, doi:10.1111/jfb.12899 A NEW COPTODON SPECIES IN WEST AFRICA? 3 of these species from remote locations over their distribution range. To recover the species diversity of this tribe and their phylogenetic relationships, a meta-analysis was performed including representative species of the different tribes and clades identified in the African cichlid phylogeny of Schwarzer et al. (2009) and Dunz & Schliewen (2013), and all cytochrome oxidase I (COI) and control region (CR) sequences labelled as C. guineensis, C. rendalli or C. zillii on GenBank. MATERIALS AND METHODS SAMPLING Samples were collected at two geographic scales: one local to investigate the genetic and morphometric diversity of cichlids in the PNBA and one continental to highlight the phylo- genetic relationships among the PNBA cichlid lineages and those of West African cichlids. In the PNBA, fish sampling was carried out by Imraguen fishermen using fixed gillnets at three locations: R’Gueiba (19∘ 25′ 27⋅1′′ N; 16∘ 27′ 56⋅2′′ W), Teichott (19∘ 32′ 38⋅2′′ N; 16∘ 24′ 37⋅6′′ W) and Iwik (19∘ 53′ 08⋅5′′ N; 16∘ 17′ 43⋅0′′ W). Each of the 393 fish collected were measured, weighed and photographed. Soft dorsal fins were clipped and stored individually in tubes of 70% ethanol. The large-scale sampling targeted some widespread species belonging to Coptodonini (C. guineensis,